Michael Whitlock. #?:Joh. ~esearch Interests. Page 1 of; ubc. ca

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v1ichael Whitlock Michael Whitlock Page 1 of; #?:Joh thitlock@zoology. ubc. ca \.ssociate Professor )ep-~lrtment of Zoology Jniversity of British Columbia 3.S., Baylor; Ph.D., Vanderbilt ~uce Scholar, World Wide Fund for Nature (WWF) - Indonesia (1990-91); -.JIHPost-doctoral Fellow, University of Chicago (1991-92); >ost-doctoral Fellow, University of Edinburgh (1992-1995) ~esearch Interests viy research focuses on evolution in structured populations: What are the forces which control the nature and distributior )f genotypes in subdivided populations and how does this affect the outcome of other evolutionary processes? These luestions and others are addressed in a variety of ways, from theoretical analyses to experimental lab model systems to ield research. have investigated the effects of non-equilibrium population structure in the partitioning of genetic variance among >opulations. Most models of population structure assume uniform populations at equilibrium, which is most unlike most tatural populations. My models have included the effects of extinctions, colonizations, population fission events, mequal population sizes, variable migration rates, and other realistic modifications of the theory. :urrently I am investigating a broad array of questions about how evolution is affected by these more generalized modes )f population structure. These questions include: What is the probability of fixing an allele? What is the load due to leleterious mutations? Is this load constant across populations? How is genetic variance maintained in structured )opulations? N' e also are concerned with the effects of small population size on the evolutionary process. Weare measuring the ~hanges in phenotypic and genetic variance as a consequence of population bottlenecks, using Drosophila melanogaster )opulations. Furthermore, we are conducting a series of experiments about the evolution of reproductive isolation in :mall populations. Using a combination of theory and experiment, we are investigating the effect of beneficial mutations m the change in fitness in small populations. am also working on many other evolutionary questions, including the evolution and measurement of phenotypic )lasticity, the effect of epistatic interactions on evolutionary change, the evolution of ploidy in finite populations, ~ffective population size, and others. ;ee a list of recent publications. illdress: Department of Zoology University of British Columbia 6270 University Blvd. Vancouver BC V6T 1Z4, Canada. Phone: (604) 822-2069 FAX: (604) 822-2416 tttp:llwww.zoology.ubc.ca/~whitlocklhp.html 2/241200:

viichael Whitlock Page 2 of: IBack to zoology faculty listing.!evolutionary Biology at UBC. lttp://www.zoology.ubc.ca/~whitlock/hp.html 2/24/200:

Michael Whitlock - Publications http://www.bcu.ubc.cai-whitiockipubjications.html Michael Whitlock Recent Publications Whitlock, M. C., and D. E. McCauley. 1990. Some population genetic consequences of colony formation and extinction: Genetic correlations within founding groups. Evolution 44: 1717-1724. Whitlock, M. C. 1992. Nonequilibrium population structure in forked fungus beetles: Extinction, colonization, and the genetic variance among populations. The American Naturalist 139:952-970. Whitlock, M. C. 1992. Temporal fluctuations in demographic parameters and the genetic variance among populations. Evolution 46:608-615. Whitlock, M. 1993. Lack of correlation between heterozygosity and fitness in forked fungus beetles. Heredity 70:574-581. Whitlock, M. c., P. C. Phillips, and M. J. Wade. 1993. Gene interaction affects the additive genetic variance in subdivided populations with migration and extinction. Evolution 47: 1758-1769. Whitlock, M. C. 1994. Fission and the genetic variance among populations: the changing demography offorked fungus beetle populations. The American Naturalist 143:820-829. Fowler, K., and M. C. Whitlock. 1994. Fluctuating asymmetry does not increase with moderate inbreeding in Drosophila melanogaster. Heredity 73:373-376. Whitlock, M. C., and M. J. Wade. 1995. Speciation: Founder events and their effects on X-linked and autosomal genes. The American Naturalist 145:676-685. Whitlock, M. C. 1995. Variance Induced Peak Shifts. Evolution 49:252-259. Whitlock, M. C. 1995. Two-locus drift with sex chromosomes: The partitioning and conversion of variance in subdivided populations. Theoretical Population Biology 48:44-64. Whitlock, M. c., P. C. Phillips, F. B. G. Moore, and S. Tonsor. 1995. Multiple Fitness Peaks and Epistasis. Annual Review of Ecology and Systematics 26:601-29. Whitlock, M. C. 1996. The Red Queen versus the Jack-of-All-Trades: evolution of specialization. The American Naturalist 148:S65-S77.. Evolutionary rates and the Whitlock, M. C., and K. Fowler. 1996 The variance among populations in phenotypic variance with inbreeding. Evolution 50:1919-1926. 10f5

Michael Whitlock - Publications http://www.bcu.ubc.ca/-whitlock/publications.html Whitlock, M. 1996. The heritability of fluctuating asymmetry and the genetic control of developmental stability. Proceedings of the Royal Society, Series B 263:849-853. Barton, N. H., and M. C. Whitlock. 1997. The evolution of meta populations. pp. 183-210 in 1. Hanski and M. Gilpin, eds., Metapopulation Biology: Ecology, Genetics, and Evolution. Academic Press. Whitlock, M. C., and K. Fowler. Evolutionary Biology 10:63-67. 1997. The instability of studies of instability. Journal of Whitlock, M. c., and N. H. Barton. 146:427-441. 1997. The effective size ofa subdivided population. Genetics Otto, S. P., and M. C. Whitlock. 1997. Fixation of beneficial mutations in a population of changing size. Genetics 146:723-733. Whitlock, M. C. 1997. Founder effects and peak shifts without genetic drift: Adaptive peak shifts occur easily when environments fluctuate slightly. Evolution 51:1044-1048. Whitlock, M. 1998. The repeatability of fluctuating asymmetry: A revision and extension. The Proceedings of the Royal Society, Series B. 265: 1429-1431. Rundle, H. D., A. 0. Mooers, and M. C. Whitlock. 1998. Single founder-flush events and the evolution of reproductive isolation. Evolution 52: 1850-1855. Whitlock, M. c., and D. E. McCauley. 1999. Indirect measures of gene flow and migration: FST doesn't equal 1/(4Nm+ 1). Heredity 82: 117-125. Whitlock, M. C., and K. Fowler. 1999. The changes in genetic and environmental variance with inbreeding in Drosophila melanogaster. Genetics 152:345-353. Mooers, A. 0., H. D. Rundle, and M. C. Whitlock. 1999. The effects of selection and bottlenecks on male mating success in peripheral isolates. American Naturalist 153: 437-444. Fowler, K., and M. C. Whitlock. 1999. The distribution of phenotypic variance with inbreeding. Evolution 53 :83-96. Whitlock, M. C., and P. C. Phillips. 1999. Genetic Drift. Encyclopedia of Life Sciences, in press. Rundle, H. D., A. 0. Mooers, and M. C. Whitlock. 1999. Experimental tests of founder-flush: A reply to Templeton. Evolution 53:1632-1633. Whitlock, M. C. and S. P. Otto. 1999. The panda and the phage: Compensatory mutations and the persistence of small populations. Trends in Evolution and Ecology 14:293-294. Whitlock, M. C. 1999. Neutral additive genetic variance in a metapopulation. 74:215-221. Genetical Research 20f5

Michael Whitlock - Publications http://www.bcu.ubc.ca/-whitlock/publications.html Phillips, P. C., S. P. Otto, and M. C. Whitlock. 2000. Beyond the average: The evolutionary importance of epistasis and the variability of epistatic effects. In Wolf, J, E. D. Brodie Ill, and M. J. Wade, (eds.) Epistasis and the Evolutionary Process. Oxford Press, Oxford. Whitlock, M. C. 2001. Dispersal and the genetic properties of meta populations. Pp. 273-282 in Clobert, J., and E. Danchin, (eds.) Dispersal. Oxford University Press, Oxford. Fowler, K., and M. C. Whitlock. 1999. The variance in inbreeding depression and the recovery of fitness in bottlenecked populations. The Proceedings of the Royal Society, Series B, 266:2061-2066. Whitlock, M. C., P. K. Ingvarsson, and T. Hatfield. 2000. Local drift load and the heterosis of interconnected populations. Heredity 84:452-457. Ingvarsson, P. K., and M. C. Whitlock. 2000. Heterosis increases the effective migration rate. Proceedings of the Royal Society 267: 1321-1326. Whitlock, M. C., and P. C. Phillips. 2000. The exquisite corpse: a shifting view ofthe shifting balance. Trends in Evolution and Ecology 15:347-348. Whitlock, M. C., and D. Bourguet. 2000. Factors affecting the genetic load in Drosophila: Synergistic epistasis and correlations among fitness components. Evolution 54: 1654-1660. Whitlock, M. C. 2000. Fixation of new alleles and the extinction of small populations: Drift load, beneficial alleles, and sexual selection. In press, Evolution 54: 1855-1861. Rundle, H. D., and M. C. Whitlock. 2001. A genetic interpretation Evolution 55: 198-201. of ecological isolation. Whitlock, M. C. 2001. Effective population size. Encyclopedia Publishing Group, London. of the Human Genome. Nature Whitlock, M. C. 2001. Genetic drift. Encyclopedia Whitlock, M. C. 2001. Inbreeding. Encyclopedia of Evolution. Oxford University Press, Oxford. of Evolution. Oxford University Press, Oxford. Whitlock, M. C., and Y. Michalakis. 2001. Metapopulations. Encyclopedia of Evolution. Oxford University Press, Oxford. Whitlock, M. C. 2001. Shifting balance. Encyclopedia Oxford. of Evolution. Oxford University Press, Phillips, P. C., M. C. Whitlock, and K. Fowler. 2001. The shape of the genetic covariance matrix changes with inbreeding. Genetics. 158: 1137-1145. 30f5

MichaelWhitlock Publications Otto, S. P. and M. C. Whitlock. 2001. Fixation times and probabilities. Encyclopedia Human Genome. Nature Publishing Group, London. http://www.bcu.ubc.ca/-whitlock/publications.html of the )t4no/~ Ives, A. R., and M. C. Whitlock. 2002. Perspective: On inbreeding and metapopulations. 295: 454-455. Science Whitlock, M. C. 2002. Selection, load, and inbreeding depression in a large metapopulation. Genetics. 160: 1191-1202. Fowler, K., and M. C. Whitlock. 2002. Environmental stress, inbreeding, and the nature of phenotypic and genetic variance in Drosophila melanogaster. Proceedings of the Royal Society, Series B. 269:677-683. M. C. Whitlock, Phillips, P. c., and K. Fowler. 2002. Persistence of changes in the genetic covariance matrix after a bottleneck. Evolution 56:1968-1975. Whitlock, M. C., and R. Burger. Still in press. The good, the bad, and the ugly: Fixation of new mutations in small populations. In Ferriere, R., U. Dieckmann, and D. Couvet, (eds.) Evolutionary Conservation Biology: The qenetics of Rarity. Oxford Press, Oxford. Whitlock, M. C. Fixation probability and time in a metapopulation. In press, Genetics. Wang, J., and M. C. Whitlock. Estimating effective population size and migration rates ITom genetic samples over space and time. In press, Genetics. Fowler, K., and M. C. Whitlock. An experimental approach to fluctuating asymmetry: Inbreeding, environmental stress, heritability and fitness. Submitted to Journal of Evolutionary Biology. Bourguet, D., J. Gair, M. Mattice, and M. C. Whitlock. Genetic recombination and adaptation to fluctuating environments: Selection for geotaxis in Drosophila melanogaster. Submitted to Heredity. Whitlock, M. C., C. K. Griswold, and A. D. Peters. The critical effective size of a population with deleterious and compensatory mutations. Submitted to Annales Zoologici Fennici. Griswold, C. K., and Whitlock, M. C. The genetics of adaptation: The roles of pleiotropy, stabilizing selection and drift in shaping the distribution of bidirectional fixed mutational effects. Submitted to Genetics. J. A. G.M. de Visser, J. Hermisson, G. P. Wagner, L. W. Ancel, H. Bagheri, J. L. Blanchard, L. Chao, J. M. Cheverud, S. F. Elena, W. Fontana, G. Gibson, T. F. Hansen, D. Krakauer, R. C. Lewontin, C. Oma, Sean H. Rice, G. von Dassow, A. Wagner, and M. C. Whitlock. Perspective: Evolution and detection of genetic robustness. To be submitted to Evolution. 40f5